The Stoor Hobbit of Guangdong: Goniurosaurus gollum sp. nov., a cave-dwelling Leopard Gecko (Squamata, Eublepharidae) from South China

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Zookeys 991: 137—153 (2020)

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The Stoor Hobbit of Guangdong:
Goniurosaurus gollum sp. nov., a cave-dwelling
Leopard Gecko (Squamata, Eublepharidae)
from South China

Shuo Qi!”, Jian Wang!", L. Lee Grismer’, Hong-Hui Chen!,
Zhi-Tong Lyu', Ying-Yong Wang'

I State Key Laboratory of Biocontrol/ The Museum of Biology, School of Life Sciences, Sun Yat-sen University,
Guangzhou, Guangdong 510275, China 2. Herpetology Laboratory, Department of Biology, La Sierra Univer-
sity, Riverside, California 92515, USA

Corresponding author: Ying-Yong Wang (wangyy@mail.sysu.edu.cn)

Academic editor: T. Ziegler | Received 31 May 2020 | Accepted 10 September 2020 | Published 11 November 2020
Attp://z00bank. org/2D9IEEFCO-B43E-4AC3-86E7-89944E54169B

Citation: Qi S, Wang J, Grismer LL, Chen H-H, Lyu Z-T, Wang Y-Y (2020) The Stoor Hobbit of Guangdong:
Goniurosaurus gollum sp. nov., a cave-dwelling Leopard Gecko (Squamata, Eublepharidae) from South China. ZooKeys
991: 137-153. https://doi.org/10.3897/zookeys.99 1.54935

Abstract

A new species of the genus Goniurosaurus is described based on three specimens collected from a limestone
cave in Huaiji County, Guangdong Province, China. Based on molecular phylogenetic analyses, the new
species is nested within the Goniurosaurus yingdeensis species group. However, morphological analyses
cannot ascribe it to any known species of that group. It is distinguished from the other three species in the
group by a combination of the following characters: scales around midbody 121-128; dorsal tubercle rows
at midbody 16-17; presence of 10-11 precloacal pores in males, and absent in females; nuchal loop and
body bands immaculate, without black spots; iris orange, gradually darker on both sides. The discovery
of yet another limestone-adapted species of Goniurosaurus in Guangdong Province underscores a growing
body of evidence for the high biodiversity of limestone habitats and brings into sharp focus the urgent

need for their conservation.

Keywords

Goniurosaurus gollum sp. nov., karst, limestone cave, taxonomy, systematics

* Contributed equally as the first authors.

Copyright Shuo Qi et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

138 Shuo Qi etal. / ZooKeys 991: 137-153 (2020)

Introduction

The eublepharid genus Goniurosaurus Barbour, 1908, currently contains 22 species
that are scattered throughout much East and northern Southeast Asia (Uetz et al.
2020; Zhu et al. 2020; Qi et al. 2020). All species of Goniurosaurus have generally
restricted, circumscribed ranges and many are restricted to habitats with either gran-
ite or limestone (karst) rock (Wang et al. 2014; Liang et al. 2018; Qi et al. 2020).
Recent molecular phylogenetic analyses resulted in the partitioning of Goniurosaurus
into four monophyletic species groups (Liang et al. 2018 ; Qi et al. 2020; Zhu et al.
2020), namely the G. kuroiwae, G. lichtenfelderi, G. luii and, G. yingdeensis groups.
The morphological comparisons also sustain the recognition of the G. kuroiwae and
G. yingdeensis groups, however, the definitions of the G. lichtenfelderi and G. luii groups
still require clarification (Qi et al. 2020).

Currently, three narrowly distributed species within the Goniurosaurus
yingdeensis group are known from the karst environments of northern Guangdong,
China: G. yingdeensis Wang, Yang & Cui, 2010, G. zhelongi Wang, Jin, Li & Gris-
mer, 2014 and G. varius Qi, Grismer, Lyu, Zhang, Li & Wang, 2020. These three
species share consistent morphological features that differentiate them from spe-
cies of the other groups: (1) base of claws sheathed by four scales, two lateral
scales of claw short and shell-shaped; (2) precloacal pores fewer than 15 in males
and absent in most females (precloacal pores present in females in G. yingdeensis
only); precloacal pores form a continuous transverse series not extending onto the
femora; (3) enlarged row of supraorbital tubercles indistinct or absent; (4) nuchal
loop rounded posteriorly; and (5) four body bands between the nuchal loop and
the caudal constriction.

During our herpetological surveys in Guangdong Province, China, three speci-
mens of Goniurosaurus were collected from a new locality during May 2020. Mor-
phological and molecular analyses place this new population within the Goniurosaurus
yingdeensis group but cannot ascribe them to any of the known species. We therefore
describe this population as a new species.

Materials and methods

Sampling

Three specimens of the undescribed species were collected from Huaiji County,
Guangdong Province, China. Following euthanasia, all specimens were fixed in 10%
formalin and transferred to 75% alcohol. Tissue samples were preserved in 99% alco-
hol and stored at -40 °C. All specimens are deposited in The Museum of Biology, Sun
Yat-sen University (SYS). Sequences of other species of Goniurosaurus generated by
Qi et al. (2020) were accessed from GenBank.

A new species of Goniurosaurus (Squamata, Eublepharidae) 139

Morphological characters

Measurements were taken following Ziegler et al. (2008) using digital calipers (Neiko
01407A Stainless Steel 6-Inch Digital Caliper, USA) to the nearest 0.1 mm. Abbreviations
of morphological characters are as follows: SVL snout-vent length (from tip of snout to
vent); TaL tail length (from vent to tip of tail); HL head length (from tip of snout to poste-
rior margin of ear opening); HW maximum head width; SE snout-to-eye distance (meas-
ured from tip of snout to the boney anterior margin of the orbit); EE eye-to-ear distance
(from the boney posterior margin of the orbit to posterior margin of ear opening); SPL su-
pralabials; IFL infralabials; N nasal scales surrounding nare; IN internasals; PostIN granular
scales bordering the internasals; PM postmentals; GP gular scales bordering postmentals;
CIL eyelid fringe scales or ciliaria,; PO preorbital scales (number of scales in a line from
posterior margin of external naris to anterior margin of the boney orbit); GST granular
scales surrounding dorsal tubercles; PTL paravertebral tubercles between limb insertions;
DTR longitudinal dorsal tubercle rows at midbody; MB scales around midbody; PP pre-
cloacal pores; PAT postcloacal tubercles. Bilateral scale counts are given as left/right.
Morphological character data of known congeners were taken from the literature
(Grismer et al. 1999, 2002; Orlov et al. 2008; Ziegler et al. 2008; Blair et al. 2009;
Wang et al. 2010, 2013, 2014; Yang and Chan 2015; Zhou et al. 2018; Qi et al. 2020;
Zhu et al. 2020) and from 34 examined museum specimens of the eight species listed

in the Appendix.

DNA Extraction, Polymerase Chain Reaction (PCR), and sequencing

Genomic DNA was extracted from muscle tissue samples, using a DNA extraction kit
from Tiangen Biotech (Beijing) Co., Ltd. Partial segments of the mitochondrial genes
16S ribosomal RNA gene (16S) and Cytochrome b gene (Cytb) were amplified. Prim-
ers used for 16S were r16S-5L (5’°- GGTMMYGCCTGCCCAGTG -3’) and 16sbr-H
(5’°- CCGGTCTGAACTCAGATCACGT-3’) (Palumbi et al. 1991) and for Cytb the
primers were L14731 (5’°- T€GTCTGAAAAACCAT TGTTG-3’) (Honda et al. 2014)
and H15149m (5’- GCMCCTCAGAAKGATATTTGYCCTCA-3’) (Chambers and
MacAvoy 1999). The PCR procedure was performed with an initial denaturation at
94 °C for 5 min, 35 cycles of 94 °C for 30 s, 55 °C for 30 s and 72 °C for 1 min, fol-
lowed by a final extension at 72 °C for 10 min (Liang et al. 2018). PCR products were
purified with spin columns and then sequenced with forward primers using BigDye
Terminator Cycle Sequencing Kit as per the guidelines on an ABI Prism 3730 auto-
mated DNA sequencer by Shanghai Majorbio Bio-pharm Technology Co., Ltd.

Phylogenetic analysis

Fifty-one sequences from 13 known Goniurosaurus species plus one outgroup sequence
from the eublepharid Hemitheconyx taylori Parker, 1930, which was used to root the

140

Shuo Qi et al. / ZooKeys 991: 137-153 (2020)

Table 1. Localities, voucher information, and GenBank accession numbers for all specimens used in this study.

Species name Locality Specimen voucher 16S Cytb References
Ingroup: Goniurosaurus
Goniurosaurus yingdeensis species group
(1) Goniurosaurus gollum sp. nov. Huaiji, Guangdong, China SYS 1002420 MT995784 MT995787 ‘This study
(2) Goniurosaurus gollum sp. nov. Huaiji, Guangdong, China SYS 1002421 MT995785 MT995788 ‘This study
(3) Goniurosaurus gollum sp. nov. Huaiji, Guangdong, China SYS 1002422 MT995786 MT995789 ‘This study
(4) G. varius Yangshan, Guangdong, China — SYS 1002330 MT995753. MT995768 Qi et al. 2020
(5) G. varius Yangshan, Guangdong, China — SYS 1002331 MT995754 MT995769 Qi et al. 2020
(6) G. varius Yangshan, Guangdong, China — SYS 1002333 MT995755 MT995770 Qi et al. 2020
(7) G. varius Yangshan, Guangdong, China — SYS 1002362 MT995756 MT995771 Qi et al. 2020
(8) G. varius Yangshan, Guangdong, China — SYS 1002363 MT995757. = MT995772 Qi et al. 2020
(9) G. yingdeensis Yingde, Guangdong, China SYS 1001271 MT995759 MT995774 Qi et al. 2020
(10) G. yingdeensis Yingde, Guangdong, China SYS 1001272 MT995760 = MT995775 Qi et al. 2020
(11) G. yingdeensis Yingde, Guangdong, China SYS 1001493 MT995761 MT995776 Qi et al. 2020
(12) G. yingdeensis Yingde, Guangdong, China SYS 1002115 MT995762 MT995777 Qi et al. 2020
(13) G. zhelongi Yingde, Guangdong, China SYS 1000816 KJ423105 =MT995778 Wang et al. 2014;

Qi et al. 2020
(14) G. zhelongi Yingde, Guangdong, China SYS 1001491 MT995763 MT995779 Qi et al. 2020
(15) G. zhelongi Yingde, Guangdong, China SYS 1001492 MT995764 MT995780 Qi et al. 2020
(16) G. zhelongi Yingde, Guangdong, China SYS 1002108 MT995765 MT995781 Qi et al. 2020
Goniurosaurus luii species group
(17) G. huuliensis Vietnam N/A AB853453 = AB853479 ~—- Honda et al. 2014
(18) G. Liboensis Libo, Guizhou, China SYS 1000217 KC900230 N/A Wang et al. 2013
(19) G. lui Jingxi, Guangxi, China SYS 1000255 KC765083 N/A Wang et al. 2013
(20) G. luii Jingxi, Guangxi, China SYS 1000256 KC765084 N/A Wang et al. 2013
(21) G. lui Cao Bang, Vietnam ZEMK 87057 EU499391 N/A Ziegler et al. 2008
Goniurosaurus lichtenfelderi species group
(22) G. bawanglingensis Bawangling, Hainan, China SYS 1002162 MT995758 MT995773 Qi et al. 2020
(23) G. bawanglingensis Bawangling, Hainan, China BL-RBZ-021 MH247190 MH247201 Liang et al. 2018
(24) G. hainanensis Jianfengling, Hainan, China SYS 1000349 KC765080 N/A Wang et al. 2013
(25) G. zhoui Central area, Hainan, China SYS 1002213 MT995766 MT995782 Qi et al. 2020
(26) G. zhoui Central area, Hainan, China SYS 1002214 MT995767 + MT995783 Qi et al. 2020
(27) G. zhoui Central area, Hainan, China BL-RBZ-001 MH247196 MH247207 Liang et al. 2018
Goniurosaurus kuroiwae species group
(28) G. kuroiwae Northern Okinawajima N/A AB853448 AB853473 ~—- Honda et al. 2014
Island, Japan

(29) G. orientalis Iejima Island, Japan N/A AB853446 AB853467 ~ Hondaet al. 2014
(30) G. splendens Tokunoshima Island, Japan N/A AB853451 AB853477 ~— Honda et al. 2014
(31) G. yamashinae Kumejima Island, Japan N/A AB853442 AB853460 Honda et al. 2014
Outgroup
(32) Hemitheconyx taylori East Africa N/A AB308457 N/A Jonniaux and

Kumazawa 2008

tree, were obtained from GenBank and composed the dataset (Table 1). DNA se-
quences were aligned by the Clustal W with default parameters (Thompson et al.
1997) and trimmed with gaps partially deleted in MEGA 6 (Tamura et al. 2013).
Two gene segments, with 486 base pairs (bp) of 16S and 396 bp of Cytb, were con-
catenated seriatim into an 882 bp sequence, and further divided into two partitions
based upon each gene. The partitions were tested in jmodeltest v2.1.2 with Akaike
and Bayesian information criteria, all resulting the best-fitting nucleotide substitution
models of GIR+I+G. Sequence data were analyzed using Bayesian inference (BI) in

A new species of Goniurosaurus (Squamata, Eublepharidae) 141

MrBayes 3.2.4 (Ronquist et al. 2012), and maximum likelihood (ML) in RaxmlGUI
1.3 (Silvestro and Michalak 2012). Two independent runs were conducted in the BI
analysis with 10,000,000 generations each and sampled every 1000 generations with
the first 25% of samples discarded as burn-in, resulting in a potential scale reduction
factor (PSRF) of < 0.005. In the ML analysis, a bootstrap consensus tree inferred
from 1000 replicates was generated. Nodes with Bayesian posterior probabilities (BPP)
>0.95 and ML support values of >70 were considered strongly supported (Huelsen-
beck et al. 2001; Wilcox et al. 2002). Uncorrected pairwise sequence divergences uti-
lizing the 16s gene were calculated using MEGA 6.

Results

The ML and BI analyses resulted in identical topologies (Fig. 1) and maintained a
high degree of consistency with recent molecular phylogenetic analyses (Liang et al.
2018; Qi et al. 2020). Uncorrected pairwise sequence divergences are reported in
Table 2. The phylogenetic analyses support previous analyses indicating that Goni-
urosaurus can be divided into four strongly supported clades, namely the G. kuroiwae
group, G. lichtenfelderi group, G. luii group and the G. yingdeensis group. The Goni-
urosaurus yingdeensis group is composed of four species with genetic differences among
them ranging from 5.5—5.9%. Three of the species are G. varius, G. yingdeensis and
G. zhelongi and the fourth species is composed of the individuals of the new popula-
tion from Guangdong Province. All specimens were recovered as monophyletic with
a high nodal support (1.00 in BI and 100 in ML) and low intrapopulational genetic
differentiation (0.0—-0.5%; Table 2). But due to low nodal support, the phylogenetic
relationships among G. varius, G. yingdeensis and G. zhelongi are still unclear. Addi-
tionally, the Guangdong population has a combination of morphological characteris-
tics distinguishing it from other species in the G. yingdeensis group as well as showing
significant morphological differences from all known congeners. Thus, we hereby de-
scribe the specimens from Guangdong Province as a new species.

Goniurosaurus gollum Qi, Wang, Grismer, Lyu & Wang, sp. nov.
http://zoobank.org/C1369EA0-37AB-457C-89C8-E98ED643E 1 E4
Figs 2, 3, 4A, 5

Material examined. Holotype. SYS 1002420, adult male (Figs 2, 3A, 4A, 5), collected
by Shuo Qi, Jian Wang and Hong-Hui Chen on 21 May 2020 from Huaiji County,
Zhaoging City, Guangdong Province, China. Exact locality available to only qualified
researchers upon request. Paratypes. One adult male (SYS 1002421) and one adult
female (SYS 1002422) share the same collection information as the holotype.
Diagnosis. Goniurosaurus gollum sp. nov. can be distinguished from all other congeners
by the following combination of characters: (1) adult body size moderate, 91.0-93.4 mm

142

| Goniurosaurus yingdeensis group

| Goniurosaurus lichtenfelderi group

0.95/50

| Goniurosaurus luii group

Goniurosaurus kuroiwae group

Ingroup

Outgroup

0.69/74
BS

1,0/94)

iosit~ SYS 1001491 (Topotype)

Shuo Qi etal. / ZooKeys 991: 137-153 (2020)

SYS r002330 (Paratype)

a SYS 1002331 (Paratype)

t SYS 1002333 (Holotype)
SYS 1002362 (Paratype)

SYS 1002363 (Paratype)

SYS r000816 (Paratype)
SYS 1001492 (Topotype)

1

Goniurosaurus varius

Goniurosaurus zhelongi

SYS 1002108 (Topotype)

SYS 1001271 (Topotype)
SYS 1002115 (Topotype)
SYS.r001272 (Topotype)
SYS 1001493 (Topotype)

Cae SYS 1002420 (Holotype)
: SYS r002421 (Paratype)
SYS r002422 (Paratype)

SYS 1002213
SYS 1002214
BL-RBZ-001 (Paratype)

1.0100} SY$ 1002162 (Topotype)
BL-RBZ-021 (Topotype)
asso yf oY 2 1000255 s ue
9 TISYS 1000256 | Goniurosaurus luii
ronoo}-ZFMK 87057 1

N/A Goniurosaurus huuliensis

N/A. Hemitheconyxtaylori -

Goniurosaurus yingdeensis

Goniurosaurus zhoui

sys r000349 Goniurosaurus hainanensis

SYS 1000217 (Paratype) Goniurosaurus liboensis
N/A Goniurosaurus kuroiwae

N/A Goniurosaurus orientalis
N/A Goniurosaurus yamashinae
N/A Goniurosaurus splendens

Goniurosaurus gollum sp. nov.

| Goniurosaurus bawanglingensis

Goniurosaurus varius

Goniurosaurus zhelongi

Goniurosaurus yingdeensis

Gonjurosaurus gullum sp. nov.

Figure |. Bayesian inference tree of 14 species of Goniurosaurus, based on the partial DNA sequences

of the mitochondrial 16S rRNA and Cytb genes. Hemitheconyx taylori is the outgroup. Numbers before

slash indicate Bayesian posterior probabilities (BPP) and numbers after slash are bootstrap support for ML

(1000 replicates) analyses.

Table 2. Uncorrected P-distance of 16S gene among 14 Goniurosaurus species used in this study.

1-3 4-8 9-12 13-16 17
Goniurosaurus gollum 0-0
sp. nov.
G. varius 5.6 0-0.1
G. yingdeensis 5.9 4.2 0-0.3
G. zhelongi 5.5 3.3 4.7 0-0.2
G. huuliensis 17.4 12.9 15.5 145 /
G. liboensis 1:7 ate loa) GIS AS erst
G. luii 164 139 145 147 14
G. bawanglingensis 17:3. 25.9" G8 158° 174
G. hainanensis 173 166 166 17.9 148
G. zhoui 16.7 163 181 17.3 16.1
G. kuroiwae 22.7. 21.2 20.33 224 22.2
G. orientalis 21.3 18.5 182 196 21.3
G. splendens PAO V7 B19,0y DNB, 22:9
G. yamashinae 22 185 188 19.9 214

18

19-21 22-23 24 25-27 28 29 30 31

/

3.6 |
hea!
BY el" Fe aE

SVL; (2) nasal scales surrounding nares seven or eight; (4) internasal single; (5) eyelid fringe
scales 59-63; (6) granular scales of upper eyelids similar in size to those on top of head; (7)
scales around midbody 121-128; (8) longitudinal dorsal tubercle rows at midbody 16 or
17; (9) paravertebral tubercles between limb insertions 25 or 26; (10) claws sheathed by

A new species of Goniurosaurus (Squamata, Eublepharidae) 143

Figure 2. A The general aspect of the adult male holotype of Goniurosaurus gollum sp. nov. (SYS 1002420)

in life B scalation and coloration characters of the head of the holotype. Photographs by Shuo Qi.

four scales, dorsal claw scales small, two lateral claw scales short and shell-shaped; (11) axil-
lary pockets deep; (12) presence of 10 or 11 precloacal pores in males and absent in females;
(13) dorsal ground color of head, body, and limbs in adults yellowish brown and mottled
with irregularly shaped dark-brown blotches; (14) nuchal loop complete, rounded posteri-
orly; (15) presence of three or four thin dorsal body bands between nuchal loop and caudal
constriction, with black anterior and posterior borders, bands usually irregularly shaped;
(16) iris orange, gradually darker on both sides.

Comparisons. Goniurosaurus gollum sp. nov. can be distinguished from the other
21 known species in the genus by the following combination of characters: base of

144 Shuo Qiet al. / ZooKeys 991: 137-153 (2020)

x 4 ; ~a i gif : ; : ,
gee SP 2mm LO, 2mm Mie Se
Figure 3. Type series of Goniurosaurus gollum sp. nov. A holotype, male, SYS r002420 B paratype, male,
SYS 1002421 C paratype, female, SYS 1002421; (1) dorsal view; (2) dorsal view of head; (3) ventral view
of head; (4) close-up of the precloacal region, the Arabic number refer to the number of precloacal pores.

Photographs by Shuo Qi.

claws being sheathed by four scales, two lateral claw scales short and shell-shaped (vs.
claws sheathed by four scales, two lateral scales of claw long, curved in G. lichtenfelderi
group and G. /uii group, and not sheathed in G. kuroiwae group); having 10 or 11 pre-
cloacal pores in males (vs. 17—46 in G. lichtenfelderi group, 16-33 in G. /uii group and
absent in G. kuroiwae group); and lacking an enlarged row of supraorbital tubercles
(present in G. lichtenfelderi group and G. /uii group).

Goniurosaurus gollum sp. nov. can be distinguished from its closest relatives in
the Goniurosaurus yingdeensis group by the following combination of characters: scales
around midbody 121-128 (vs. 101-110 in G. varius, 101-116 in G. yingdeensis, 99-
109 in G. zhelongi); longitudinal dorsal tubercle rows at midbody 16 or 17 (vs. 21-24
in G. varius, 20-25 in G. yingdeensis, 23-28 in G. zhelongi); absence of ten precloa-
cal pores in females (vs. present in G. yingdeensis); nuchal loop and body bands im-

A new species of Goniurosaurus (Squamata, Eublepharidae) 145

a:
Ay. Bes setae

Figure 4. Comparisons of iris color with three closely related congeners A Goniurosaurus gollum sp. nov.
(holotype, SYS 1002420) B Goniurosaurus varius (holotype, SYS 1002333) © Goniurosaurus yingdeensis
(holotype SYSr000504) D Goniurosaurus zhelongi (holotype, SYS 1000770). Photographs by Shuo Qi and
Ying-Yong Wang.

maculate (vs. having black spots in G. varius); iris orange, gradually darker on both
sides (vs. iris orange-red in G. varius, iris gray and becoming orange near the pupil in
G. yingdeensis, iris gray-white and tinged with orange in G. zhelongi). Additional com-
parisons of morphological characteristics are provided in Table 4 and Figure 4.
Description of holotype. Adult male with original tail; SVL 93.4 mm; HL
24.2 mm; HW 16.2 mm; SE 9.3 mm; EE 9.3 mm; SVL:HL 3.9; HL:HW 1.5; SE:EE
1. Head triangular, wider than neck, covered with granular scales, densely interspersed
with tubercles in the temporal and occipital regions; area between orbits uniformly
covered by small granular scales; supraorbital tubercles nearly uniform in size; scales of

146 Shuo Qi et al. / ZooKeys 991: 137-153 (2020)

rostrum slightly larger than those in between orbits; rostral convex and hemic-elliptic,
1.8 times as broad as high, middorsal portion of rostral partially sutured dorsomedially,
bordered laterally by first supralabial and prenasal, dorsolaterally by supranasal, dor-
sally by one internasal; external nares oval, surrounded by 8/9 nasals each, anteriorly
by prenasal and supranasal, dorsally by supranasal and a granular scale, posteriorly by
7/8 smaller granular scales, and ventrally by the prenasal; prenasal with long recurved
ventral portion; supranasals large, separated by one internasals; supralabials rectangu-
lar, 10/10; preorbital scales 16/17; eyes relatively large, pupils vertical; eyelid fringe
scales 59/60; outer surface of upper eyelid composed of granular scales of about same
size of those on top of head; external auditory meatus circular, tympanum deeply re-
cessed; mental triangular, bordered laterally by first infralabial and posteriorly by three
postmentals; infralabials rectangular, 10/10; gular scales juxtaposed uniform granular,
abruptly into flat juxtaposed pectoral scales, and grading posteriorly imbricated larger
ventral scales. Tongue with a small notch at tip. Crowns of teeth expanded, occlusal
margins bearing multiple ridges.

Dorsal surface of neck and body covered with uniform granular scales, interspersed
with densely sharply pointed conical tubercles; scales around midbody 125; longitudi-
nal rows of dorsal tubercles at midbody 16; vertebral row of scales indistinct; paraver-
tebral tubercles between limb insertions 25; dorsal body tubercles surrounded by 9-10
granular scales; dorsal scales grading ventrally into larger flattened imbricate ventral
scales; ten precloacal pores in a transverse series; postcloacal region greatly swollen,
covered with imbricate flattened scales, containing 2/2 postcloacal tubercles laterally
at level of the vent.

Tail original, long and thin, thickest at base, bearing whorls anteriorly, gradually
narrowing to the tip; composed of nine recognizable annuli anteriorly that are 8—9
scales wide, annuli fade abruptly posteriorly into flat juxtaposed scales; incorporating
2-8 sharply pointed conical tubercles in a transverse row, tubercles do not encircle the
tail; ventral caudals larger and more nearly square than dorsal caudals.

Limbs relatively long and slender; dorsal surface covered with granular scales,
densely interspersed with tubercles; ventral surface covered by flat scales, juxtaposed,
subimbricate or imbricate; dorsal surface of pes and manus covered with granular
scales, interspersed with several conical tubercles on top of pes, lacking tubercles on
top of manus; hind limbs slightly larger than forelimbs; ventral surfaces of pes and
manus covered with large granular scales; axillary pockets deep; subdigital lamellae
wide, 10/10 on Finger I, 12/14 on Finger II, 15/16 on Finger III, 16/14 on Finger
IV, 13/14 on Finger V, 12/12 on Toe I, 15/ 16 on Toe H, 20 / 19 on Toe HI, 22 / 23
on Toe IV, and 18 / 20 on Toe V; fingers laterally compressed, relative finger lengths
I<V<II<IV<III; toes laterally compressed, third toe nearly as long as the fourth toe,
relative toe length I<II<V<HI<IV; base of claws sheathed by four scales, two lateral
scales of claw short, asymmetrical shell-shaped.

Coloration in life. Dorsal ground color of head, neck, body, and limbs yellowish
brown, mottled with irregularly shaped dark-brown blotches; nuchal loop complete and
rounded posteriorly, anterior ends terminating at posterior margins of ear openings,

A new species of Goniurosaurus (Squamata, Eublepharidae) 147

Table 3. Mensural (mm) and meristic diagnostic characters (minimum/maximum) of type series of Goni-

urosaurus gollum sp. noy. See Materials and methods for abbreviations. * holotype, # paratype.

Morphological character SYS 1002420 * SYS 1002421 # SYS 1002422 #
Sex male male female
SVL 93.4 93.3 91.0
TaL 83.5 Regenerated 76.0
HL 24.2 24.0 23.3
HW 16.2 15.3 14.8
SE 9.3 9.3 9.1
EE 9.3 9.4 8.6
SPL 10/10 10/10 10/10
IFL 10/10 10/10 10/10

N 8/9 9/8 8/9
IN 1 1 1
PostIN 2 2 2
PM 2 3 3
GP Ws 8 8
CIL 59/60 59/61 63/63
PO 16/17 15/15 16/17
GST 9-10 9-10 9-11
PTL 25 26 25
DTR 16 16 17
MB 125 121 128
PP 10 11 Absent
PAT 2 2 2

edged dorsally and ventrally by wide dark-brown margin, yellow. Only two complete
body bands can be recognized between nuchal loop and caudal constriction: first band
located posterior to axilla; second band inserts onto dorsal surface of thigh, bands on
limbs dirty yellow, lacking dark spots, edged by broad dark-brown borders anteriorly
and posteriorly, other blotches incomplete, not forming a complete bands. Supralabials
and infralabials grayish brown; pupils vertical and appear black; iris orange, gradually
darkening on both sides; dorsal surface of limbs light grayish brown with dark brown
and dirty yellow tubercles and dark spots and blotches; chin, throat, thorax, and ventral
surfaces of body pink, tinged brownish, with dark-brown lateral spots; ventral surface
of limbs pink, tinged brownish, without dark-brown spots; digits light grayish brown;
ground color of original tail dark brown with nine immaculate white caudal bands
completely encircling the tail, and a white tip. Body color becomes darker after capture.

Coloration in preservative. Dorsal ground color of head, body, and limbs become
darker; ventral surface faded to grayish white; all darker spots, blotches and bands on
dorsal surface blurred.

Variations. Measurements of type series specimens are shown in Table 3. Two
paratypes have same approximate measurements as holotype, but have significant vari-
ations in coloration. Male paratype (SYS 1002421, Fig. 3B) has four body bands, the
second band extending a three forked branch connecting with third band. Female
paratype (SYS 1002422, Fig. 3C) has three complete body bands, second band extend-
ing a branch forward to middle of body, original tail has ten immaculate white caudal
bands completely encircling tail, and dark-brown tip.

Shuo Qi etal. / ZooKeys 991: 137-153 (2020)

Figure 5. The holotype of Goniurosaurus gollum sp. nov. (SYS 1002420) at its habitat: a barren limestone

cave of Guangdong, China. Photograph by Shuo Qi.

149

A new species of Goniurosaurus (Squamata, Eublepharidae)

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Etymology. The specific epithet “gollum” is named after the fictional character,
Gollum, from J.R.R. Tolkien’s The Lord of the Rings book series. This new species and
Gollum have similar cave-dwelling habit and emaciated body. We suggest the common
name as “Gollum Leopard Gecko”, and according to the type locality, we suggest the
Chinese formal name as “guang dong jian hw” (J ARIE).

Distribution and ecology. Currently, Goniurosaurus gollum sp. nov. is known only
from Huaiji County, Guangdong Province, China. All individuals were found within
a barren limestone cave approximately 50 m from the cave entrance at night after
2130 hrs (Fig. 5). The surface of the cave is covered with bat (unidentified) and bird
(Apus pacificus) droppings. Stalactites are suspended from the roof and there is no veg-
etation. Duttaphrynus melanostictus was observed in the same area.

Discussion

With the description of Goniurosaurus gollum, there are now 23 known species in
the eublepharid genus Goniurosaurus, 15 of which occur in China. The G. yingdeensis
group is endemic in Guangdong Province, with only two species recognized previously,
but the discoveries of G. varius (Qi et al. 2020) and G. gollum adds to this and indicates
there might be more new species yet to be discovered.

Although the species of Goniurosaurus are also called “cave geckos”, they prefer in-
habiting the forest floor, limestone cliffs, rocky and cement drains along the road, and
near the entrances of caves. However, the cave-dwelling species G. gollum appears to be
a true cave dweller as opposed to other Goniurosaurus. The fact that all three specimens
were found at least 50 m inside the cave from its entrance supports this hypothesis.

These results support the growing body of evidence from China (Luo et al. 2016)
and elsewhere (Grismer et al. 2016 and references therein) that karstic habitats not only
provide a substrate for the evolution of new species—especially gekkotan lizards—but
maintain levels of biodiversity that rival that of other tropical habitats. Yet they are some
of the most imperiled and least protected habitats in the world (Day and Urich 2000;
Gillieson 2005). The discovery of yet another new species in the vast karstic landscape of
northern Guangdong underscores this fact and brings into sharp focus the urgent need
to protect these unique landscapes. We aim to protect G. gollum from the pet trade, by
withholding its precise locality. However, these data are available to qualified researchers
upon official request to The Museum of Biology, Sun Yat-sen University.

Acknowledgements

We thank Truong Nguyen, Nikolai L. Orlov, Minh Duc Le and Thomas Ziegler for their
helpful comments and suggestions on the manuscript. We thank the Zhaogqing City
Forestry Bureau for provided information. This study is supported by the Goniurosaurus
yingdeensis Population Conservation Research of Guangdong Shimentai National Nature
Reserve and the Mohamed bin Zayed Species Conservation Fund (project no.142599 12).

A new species of Goniurosaurus (Squamata, Eublepharidae) 151

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Appendix

Examined specimens

Goniurosaurus bawanglingensis (IN = 3): China: Hainan Province: Bawangling National
Nature Reserve: SYS r001075, 1670, 2162.

Goniurosaurus hainanensis (N = 2): China: Hainan Province: Jianfengling National
Forest Park: SYS r000349; Baoting County: SYS 1001270.

Goniurosaurus liboensis (N = 3): China: Guizhou Province: Libo County: Maolan
National Nature Reserve: SYS r000217, 854, 855.

Goniurosaurus luii (IN = 4): China: Guangxi Zhuang Autonomous Region: Jingxi City:
SYS 1000255, 256, 859, 860.

Goniurosaurus varius (N = 5): China: Guangdong Province: Yangshan County: SYS
1002330, 2331, 2333, 2362, 2363.

Goniurosaurus yingdeensis (IN = 10): China: Guangdong Province: Yingde City: SYS
1000501, 503, 504, 535, 536, 550, 1271, 1272, 1493, 2115.

Goniurosaurus zhelongi (N = 5): China: Guangdong Province: Yingde City: SYS
1000816, 1491, 1492, 2011, 2108.
Goniurosaurus zhoui (N = 2): China: Hainan Province: SYS 1002213, 2214.